AMARANTHUS L
54. MENISPERMACEAE
CERATOPHYLLUM L.
Ceratophyllum demersum L., Sp. PI. 992. 1753 Monoecious, submerged, aquatic herb, rootless, to about
1 m long, dark green, unbranched or branched, with a single lateral branch produced at a node. Leaves whorled, 6-12 per node, 1-1.5 cm long, dichotomously dissected once or twice, the divisions mostly filiform, minutely denticulate, the whitish teeth on a broad base of green tissue. Flowers rare, unisexual, sessile, solitary in axil of one leaf of a whorl, ca 1 mm long; staminate flowers with sepals 10-15, basally connate; corolla lacking; stamens 10-20, spirally arranged on a flat receptacle; filaments very short; anthers linear-oblong, with a thickened, produced connective. Pistillate flowers with perianth similar to staminate flowers; ovary gradually tapered to slender style; pistil longer than sepals; stigma 1, undiffer- entiated from style. Nuts also rare, axillary, tuberculate, with 2 spines at base, ca 4 mm long, tipped by the long persistent style. Croat 5738.
Occasional, in coves around the island, occurring with but much less common than Hydrilla verticillata (L.f.) Royle (16. Hydrocharitaceae). Seasonal behavior unknown.
This distinctive aquatic is not confused with any other in Panama.
Cosmopolitan distribution. In Panama, it occurs prob- ably in all lakes or slow-moving, freshwater habitats.
54. MENISPERMACEAE
Lianas or vines. Leaves alternate, petiolate; blades simple, entire or crenate (may be slightly lobed in Odontocarya);
venation palmate or blades only palmiveined at base;
stipules lacking. Flowers unisexual (dioecious), generally actinomorphic, in axillary, bracteate, often fasciculate panicles or racemes; sepals and petals 3 or 6 (1 or 4 in Cissampelos; petals lacking in Abuta), the sepals free, in 1 or 2 series; petals often connate; stamens 6 (or 4 in Cissampelos), opposite the petals; anthers 2- or 4-celled, the thecae often separated by a connective, dehiscing longitudinally; gynoecium of 1-6 free pistils; ovary su- perior, 1-locular; placentation parietal; ovules 2, aborting to 1, anatropous; style 1, very short, or the stigma sessile;
stigma lobed or incised. Fruits drupaceous; seed usually lacking endosperm.
These uncommon vines or lianas are distinguished by unisexual and usually three-parted flowers, often bearing a double whorl of sepals, leaves palmately veined (at the base), and curved seeds.
Pollination systems are unknown. Plants are both dioecious and not very common.
Species with known fruits (and certainly Cissampelos species) are probably eaten by birds. Cissampelos are also eaten by white-faced monkeys (Oppenheimer, 1968).
About 65-70 genera and 350-400 species; mostly in the tropics.
ABUTA Aubl.
Abuta panamensis (Standl.) Kruk. & Barn., Mem. New York Bot. Gard. 20:22. 1970
Hyperbaena panamensis Standl.
Dioecious liana to 35 m high or tree to 6 m tall; trunk not known, probably flattened; stems terete, the older stems lenticellate. Leaves alternate; petioles (1)4-6(9) cm long, usually curved and enlarged near apex; blades variable, elliptic to oblong or ovate, acute and apiculate or short- acuminate at apex, obtuse to rounded or truncate at base,
KEY TO THE SPECIES OF MENISPERMACEAE
Blades peltate; staminate flowers with 4 sepals and 4 stamens, the petals connate; pistillate flowers with 1 sepal and 1 petal:
Inflorescence bracts to 1.5 cm long and wide; carpels glabrous; drupes 5-7 mm long; leaves and stems sparsely pubescent, the trichomes long and spreading .... Cissampelos tropaeolifolia DC.
Inflorescence bracts usually less than 1 cm long; carpels densely pubescent; drupes 4-5 mm long; leaves and stems moderately to densely pubescent, the trichomes long and sericeous or short and tomentose Cissampelos pareira L.
Blades basifixed; staminate flowers with 6-16 sepals and 6 stamens; petals various; pistillate flowers with 6 sepals and 6 petals or petals lacking:
Blades densely white-woolly beneath Chondrodendron tomentosum R. & P.
Blades not densely white-woolly beneath:
Plants herbaceous vines; blades broadly ovate, usually conspicuously cordate, with domatia or glands at base; sepals 6; petals 6:
Blades short-pubescent at least on veins below; drupes ca 1 cm long Odontocarya tamoides (DC.) Miers var. canescens (Miers) Barn.
Blades glabrous; drupes ca 2 cm long Odontocarya truncata Standl.
Plants woody lianas; leaf blades mostly elliptic, rarely subcordate, lacking domatia or glands at base; sepals 6 in 2 series; petals lacking:
Petioles mostly more than 4 cm long; staminate inflorescences more than 10 cm long; flow- ers obviously pedicellate Abuta panamensis (Standl.) Kruk. & Barn.
Petioles usually less than 3 cm long; staminate inflorescences less than 10 cm long; flowers sessile or nearly so Abuta racemosa (Thunb.) Tr. & Planch.
Fig. 229. Nymphaea ampla
Fig. 230. Abuta racemosa
Fig. 231. Chondrodendron tomentosum
54-
MENISPERMACEAE/CISSAMPELOS393
6-24 cm long, 3-14 cm wide, glabrous above, sparsely puberulent or glabrous below; veins at base 3-5. Stami- nate inflorescences racemose, to 10-15 cm long, the branches reduced to short-stalked, 3-flowered cymes;
branches and pedicels appressed-pubescent; pedicels to 3 mm long; outer sepals 3, narrowly ovate, ca 1 mm long, the inner sepals 3, broadly ovate to obovate, 2-2.8 mm long, fleshy, granular-puberulent on inside, ± strigose on outside; petals lacking; stamens 6, 1.5-2.2 mm long;
filaments somewhat flattened, pubescent at least on one margin, connate into a column toward the base; anthers ca 0.2 mm long, dehiscent longitudinally; pistillode lacking. Pistillate inflorescences racemose, axillary, pe- dunculate or subsessile, 7-12 cm long, with few flowers;
sepals and petals similar to staminate flowers; staminodia 6, 2-2.3 mm long, short-pilose in basal half, glabrous above; ovary densely pubescent; styles 3. Fruiting pedun- cles stout, ca 2 mm long; fruits of 3 drupes, the drupes oblong, ca 2.5 cm long and 1.3 cm diam, yellow-orange, densely short-pubescent, sessile. Bangham 484, Bailey 101, Standley 28417.
A number of collections exist, but the plant has not been seen recently on the island. Seasonality not deter- mined. Flowers mostly from July to September elsewhere in Central America, with the fruits maturing from March to May. Some flowers have also been seen in November and February.
Mexico to Panama. In Panama, known from tropical moist forest in the Canal Zone and Veraguas.
Abuta racemosa (Thunb.) Tr. & Planch., Ann. Sci.
Nat. Bot, ser. 4, 17:48. 1862
Dioecious liana, growing into canopy; trunk flattened, 5-20 cm diam; stems puberulent when young, becoming glabrous, the apices often attenuated and twining. Leaves alternate; petioles 0.7-4.5 cm long, usually curved and enlarged near apex; blades oblong-elliptic to oblanceolate, acuminate to bluntly acuminate, rounded, truncate, or subcordate at base, 5-15.5 cm long, 2-8.5 cm wide, stiff, glabrous but with sparse appressed pubescence below and stiff erect trichomes on major veins above, the mar- gins entire to obscurely crenulate; veins at base 3-5, the midrib arched. Staminate inflorescences narrowly pseudo- racemose, 5-10 cm long, the branchlets reduced to short- stalked, 3-flowered cymules; pedicels to 1.7 mm long;
pedicels and sepals with dense, erect or appressed puber- ulence; outer sepals 3, narrowly triangular or lanceolate, ciliate, ca 1.3 mm long, the inner sepals 3, ovate, thick, valvate, held closely together in basal half, somewhat spreading above the middle, acute at apex, ± glabrous to granular-puberulent on inside, with appressed grayish pubescence on outside; petals lacking; stamens 6, in- cluded, slightly more than 1 mm long, in 2 series, the inner 3 somewhat larger with the filaments connate into a column and the anthers swollen, the thecae apical, widely separated by a swollen connective, the outer 3 with the filaments free and the anthers minute. Pistillate inflorescences similar to staminate but apparently shorter;
pistil 3-carpellate; ovary densely sericeous, curved; styles
apical and possibly fused to each other in flower but free at least in juvenile fruit, curved outward, pointed at apex. Fruits 3-parted, the carpels drupaceous, oblong- obovate, orange to black at maturity, 2-2.3 cm long, 1.1-1.2 cm diam, tomentulose when young, glabrate;
seeds 1 per carpel, each surrounded by a fleshy white mesocarp. Croat 15004.
Occasionally seen, though likely to be common in the canopy; the flowers are small and difficult to see. Most collections have been made in tree-fall areas. Flowers in the early rainy season (May to July), with the fruits prob- ably maturing from August to October.
Panama, Colombia, and Bolivia. In Panama, known only from tropical moist forest on BCI and in Darien.
See Fig. 230.
CHONDRODENDRON R. & P.
Chondrodendron tomentosum R. & P., Syst. Veg.
261. 1798
C. hypoleucum Standl.
Large dioecious canopy liana; trunk near the ground to ca 10 cm diam; stems terete, puberulent to tomentose (striate when dry). Petioles 4-14 cm long, puberulent to tomentose; blades ovate to suborbicular, blunt to apicu- late (rarely emarginate) at apex, truncate to cordate at base, 10-20 cm long, 9-18 cm wide, subcoriaceous, gla- brous above, white-woolly below; veins at base 5, pal- mate, with 2-4 additional pairs of major lateral veins above. Staminate inflorescences paniculate; panicles fascicled in leaf axils, often on older stems, to 10 cm long, the branches short; all axes densely tomentose;
flowers greenish-white; sepals 9-16, the outermost re- duced, tomentose or ciliolate, the inner 6-8 much larger, to 3 mm long, oblong-obovate to obovate, recurved, glabrate or inconspicuously pubescent, ciliolate; petals usually 6, minute and obscured by sepals, less than 1 mm long; stamens 6, ca 1.5 mm long; anthers ± horizon- tal, affixed subbasally, the connective produced beyond the thecae and usually directed inward. Pistillate inflores- cences and flowers similar to staminate but with gynoe- cium of 6 pistils, the pubescence dense, short-appressed;
style 1 per carpel, simple, ca 0.5 mm long, slightly curved.
Drupes oblong-ovoid, ca 1.5 mm long, narrowed at base into a short stipe, tomentose to glabrous in age. Croat 13801.
Uncommon. Known to flower in late July and early August.
Panama to Peru and Bolivia. In Panama, known from tropical moist forest on BCI and in Darien.
See Fig. 231.
CISSAMPELOS L.
Cissampelos pareira L., Sp. PL 1031. 1753 Bejuco de cerca, Alcotan
Generally a slender twining vine; stems, petioles, and lower surface of leaves loosely pubescent with long and
394
DICOTYLEDONEAEsericeous or short and puberulent trichomes, the upper leaf surface glabrate or sparsely pubescent. Leaves ob- scurely peltate; petioles 3.5-7 cm long, slender; blades broadly ovate, usually obtuse and mucronate at apex, truncate to cordate at base, 2-12 cm long and wide; vena- tion palmate. Staminate inflorescences lacy and branched many times, usually axillary, often at leafless nodes;
bracts less than 1 cm long; flowers greenish, ca 3 mm diam; sepals 4, much longer than the cone-shaped corolla;
stamens united, the apex flaring; anthers 4. Pistillate flowers fasciculate along axillary branches, the branches often several at a node, the fascicles each subtended by a small, ovate or reniform, leaflike bract; sepals and petals each 1 per flower, caducous, borne at the gibbous base of the carpel; carpel 1, densely pubescent; stigma deeply tri- lobate. Drupes suborbicular, ± flattened, red, 4-5 mm long, pubescent; seed brown, flattened, verrucose, 3-4 mm long, the embryo horseshoe-shaped. Croat 4397, 11997.
Generally abundant in clearings; rare in the forest.
Flowers and fruits throughout the year.
Throughout tropics of the world. In Panama, known from tropical moist forest in the Canal Zone, Bocas del Toro, Veraguas, Los Santos, Panama, and Darien, from premontane moist forest in the Canal Zone and Panama, from premontane wet forest in Chiriqui, Los Santos, Code, and Panama, and from tropical wet forest in Code and Darien.
Cissampelos tropaeolifolia DC, Reg. Veg. Syst. Nat.
1:532. 1818
Twining vine; stems, leaves, and inflorescence branches usually with long whitish trichomes. Leaves peltate;
petioles slender, 3-7 cm long; blades ovate to suborbic- ular, rounded to obtuse or acuminate and mucronate at apex, truncate to cordate at base, 5-11 cm long and wide, glabrous to sparsely long-pubescent above, paler and usually long-pubescent or puberulent below; venation palmate. Staminate inflorescences of fasciculate dichasia in leaf axils or on short axillary branches in the axils of reduced leaves or bracts, the bracts to 1.5 cm long and 2.5 cm wide; flowers minute; sepals 4,1-1.5 mm long, ca 1 mm wide, usually pubescent on outside; corolla greenish, campanulate, 0.5-1 mm diam, glabrous; stamens with filaments united into a tube; anthers 4, borne on filament tube, glabrous. Pistillate flowers fasciculate in the axils of bracts on secondary branches; sepal 1, ovate to obovate, ca 1 mm long, glabrous or puberulent; petal 1, greenish, suborbicular, ca 0.5 mm long and to 1 mm wide; carpel 1, gibbous, sessile, glabrous; stigma trilobate. Drupes ob- ovoid, flattened, 5-7 mm long, 4-5 mm wide, red, pubes- cent; seed verrucose, the embryo horseshoe-shaped.
Croat 9265.
Rare in the forest. Flowers and fruits throughout the year.
Southern Mexico to northern South America. In Pan- ama, known from tropical moist forest in the Canal Zone, Bocas del Toro, San Bias, Chiriqui, and Darien, from
premontane wet forest in the Canal Zone and Chiriqui, and from tropical wet forest in Darien.
ODONTOCARYA Miers
Odontocarya tamoides (DC.) Miers var. canescens (Miers) Barn., Mem. New York Bot. Gard.
20:89.1970
O. paupera (Griseb.) Diels
Slender vine; older stems lenticellate. Leaves alternate;
petioles slender, 3-8.5 cm long; blades ovate to subhas- tate, obtuse to acuminate and mucronulate at apex, usu- ally shallowly cordate at base (acute on juveniles), 3-10 cm long, 3-8.5 cm wide, glabrous above, with conspic- uous glandular areas at base of blades near petiole, pubes- cent below with short, spreading trichomes especially on veins; venation at base palmate. Inflorescences axillary or supra-axillary, solitary; rachis glabrous or puberulent;
staminate inflorescences pseudoracemose, to 22 cm long, with flowers in fascicles of 2-6, occasionally with several short branches; pedicels 3-6 mm long; sepals 6, mem- branaceous, the outer 3 ± ovate, to 1.5 mm long and 1 mm wide, the inner 3 ± oblanceolate, 2-4 mm long, 1.5-2.5 mm wide; petals 6, the outer 3 greenish, some- what fleshy, bowl-shaped, 2-2.5 mm long, 1.5-2 mm wide, the inner 3 similar but smaller; stamens 6, 1-2 mm long, the filaments connate about half their length; an- thers 6, ca 0.5 mm long. Pistillate inflorescences strictly racemose, with 6-12 loosely spaced flowers; pedicels to 1.4 cm in fruit; sepals and petals as in staminate flowers;
staminodia 6; ovary glabrous. Fruits of 1-3 drupes, ellip- soid, ca 1 cm long and 7 mm diam, reddish-orange, drying blackish. Croat 5631, 5640.
Rare in clearings and occasional in the canopy of the forest. Seasonal behavior uncertain. Known to flower and fruit from May to October.
Southern Mexico to Colombia, Venezuela, the Gui- anas, and northeastern Brazil; Lesser Antilles. In Panama, known from tropical moist forest in the Canal Zone, Panama, and Darien and from tropical dry forest in Panama.
Odontocarya truncata StandL, J. Arnold Arbor.
11:121. 1930
Glabrous, slender vine. Leaves alternate; petioles 4.5-7.5 cm long; blades ovate, acuminate, truncate to shallowly cordate at base, 8-13 cm long, 5-9 cm wide, with sunken, discolored domatia in axils on lower surface; venation palmate at base. Inflorescences racemose, axillary (some- times at leafless nodes), solitary; sepals of staminate flow- ers 6, membranaceous, the outer 3 ovate, ca 0.5 mm long and wide, the inner 3 obovate, concave, ca 2 mm long and 1.5 mm wide; petals of staminate flowers 6, narrowly oblanceolate, ca 1.5 mm long and 0.5 mm wide; stamens 6, ca 1.3 mm long; filaments ca 1 mm long, the outer 3 connate to the middle, the inner 3 connate their entire length. Pistillate infructescences 10-20 cm long; fruiting pedicels 1-1.5 cm long; pistillate flowers unknown. Fruits